Male Appendages Research Articles

Two new genera of coastal Talitridae (Amphipoda: Senticaudata) from Chile, with the first record of Platorchestia Bousfield, 1982 in the southeastern Pacific coast

Using samples of talitrid amphipods collected on the coast of Chile, we describe two new monotypic genera characterized by particular types of sexual dimorphism, and we report the first record of the genus Platorchestia Bousfield, 1982 in the southeastern Pacific coast. In the subfamily Platorchestiinae, the genus Atacamorchestia gen. nov. and its new species Atacamorchestia atacamensis sp. nov. was collected in marine rocky coasts, in several localities in northern Chile. This new taxon is characterized principally by presence of T setae in male appendages, a feature shared with the genus Vallorchestia from Australia, but with a different distribution of these structures. The first specimens of Platorchestia in the Southeastern Pacific were obtained in Antofagasta, northern Chile, however they were identified only to genus level because a more in-depth taxonomic analysis will be necessary to determine their status. Finally, in the subfamily Talitrinae, Lafkenorchestia gen. nov. and its new species Lafkenorchestia oyarzuni sp. nov. were found on rocky marine shores and in estuaries in southern Chile, between La Araucania and Los Lagos regions. This new taxon is characterized by distinct modifications in basis and merus of male pereopod 7, showing some morphological similarities with the New Zealand genera Subantarctorchestia and Pictonorchestia. For each species, descriptions and illustrations are presented.

Morphological and molecular evidence supports the species status of the Italian endemic Coenagrion castellani Roberts, 1948 (Coenagrionidae)

Coenagrion castellani Roberts, 1948 was described from Italy as a distinct species almost 75 years ago but has generally not been recognised or was treated as a subspecies of C. mercuriale (Charpentier, 1840). Populations south of the Alps were recently shown to be completely isolated genetically from those in North Africa and elsewhere in Europe. As markings and male appendages also allow for easy separation in the field, C. castellani is best treated as a good species, the 146th odonate species known from Europe and the second one that is endemic to Italy. Its identification and occurrence are reviewed. North African populations are distinct genetically too, but not in morphology. Whether these should be treated as a distinct taxon, e.g. as the subspecies C. mercuriale hermeticum (Selys, 1872), requires further research.

Morphology of Three Sphyrion (Copepoda: Siphonostomatoida: Sphyriidae) Species Infecting Teleost Fishes off South Africa with the First Description of Males of Two Species

Sphyrion is a member of the family Sphyriidae and currently consists of three valid species, namely S. laevigatum, S. lumpi and S. quadricornis. Species of Sphyrion are mesoparasites of a variety of bony fishes and occur worldwide. The bodies of the post-metamorphosis females are highly transformed with an elongated cephalothorax bearing conspicuous lateral processes. The structure of the cephalothorax is mostly used in the identification of the different species. The morphology of the adult males resembles those of the lernaeopodid males. Species of Sphyrion were collected from various fish hosts off the Indian and Atlantic Oceans off South Africa. Selected specimens were studied using dissection and compound light microscopy as well as scanning electron microscopy. Similar to previous reports, the morphology of specimens of the same species infecting the same host species is highly variable; thus, the structure of the cephalothorax should be considered together with the rest of the body features for identification. Additional information is provided regarding the morphology and armature of the appendages of the post-metamorphosis females of all three species, which resemble previous descriptions, although no detailed comparisons could be done. This is the first collection of descriptions and illustrations of the males of S. laevigatum and S. quadricornis. Male appendages of all three species are very similar with small differences in the armature. New geographical and host reports of the Sphyrion species are provided.

Sexual dimorphism and static allometry in the South African scorpion Opistophthalmus karrooensis

Sexual dimorphism in scorpions is generally retrieved for a similar subset of body parts and is presumed to result from alternative types of selection operating on homologous structures in each sex. Even so, investigations which use static allometry as a method for testing specific hypotheses relating to selection remain limited. Sexual dimorphism in the majority of South African scorpion species is poorly investigated, especially in the genus Opistophthalmus, of which the endemic Karoo burrowing scorpion O. karrooensis is nearly completely data deficient. The current study investigates patterns of sexual dimorphism and static allometry in O. karrooensis, based on external morphology. Following character scaling on carapace length, patterns of sexual dimorphism in this species indicate the elongation of male appendages (pedipalp segments, legs, pectines, metasoma, and telson), while females display wider bodies and wide, thick pedipalp segments. These patterns possibly point to an enhanced reproductive output and parental care in females, coupled to a sedentary lifestyle and generalist feeding strategy. Conversely, the male morphology is likely adapted for greater vagility, but should also function during mate-trailing and sexual interactions. In line with these explanations, the body parts in both sexes indicate static allometric patterns pointing (at least in part) to the potential influence of either natural or sexual selection, but further reveal the possibility of intense selection on a number of body parts. Taken together, the selective forces which shape scorpion morphology may be complex, relating to multiple functions of body parts and life-history differences between the sexes.

Male terminalia of Cercopidae (Hemiptera, Cicadomorpha): towards a consensus terminology

The study of male genital appendages is often necessary to identify a species and to characterise the higher systematics ranks for the Cercopidae, a large family of Hemiptera. Therefore, many authors have used them in their work but without any clear consensus on the terms used for each part constituting the male terminalia. A standardised terminology is important for the quality of a taxonomic description but even more essential when we want to compare species and establish a primary homology between states of character and their use in the frame of phylogenetic analysis. The use of a consensus terminology should ensure that we are all observing, speaking and describing the same genital appendage and comparing homologous characters. In order to propose a consensus terminology, we have reviewed all the major works on the anatomy of terminalia for the family since the first description using those characters in 1922. We proposed the use of consensual terms, listed with their definitions. In addition we studied a diversified panel of male specimens, chosen in order to represent as many Cercopidae tribes as possible. We categorised five different groups of Cercopidae according to their male terminalia structures. This opens the reflection on the evolutionary patterns for these structures.

Interspecific crossing between blue‐tailed damselflies Ischnura elegans and I. senegalensis in the laboratory

AbstractIschnura species (Odonata) are among the most common damselflies in the world, which often exhibit female color polymorphisms. One morph, called androchrome, is similar to males in its color pattern, whereas the other morphs, generally referred to as gynochromes, exhibit female‐specific colors. In several Ischnura species, the female polymorphism is heritable, although molecular and genetic mechanisms remain largely unknown. The dominant‐recessive patterns of the female color morphs may differ between species. For example, androchromic females are dominant to gynochromic females in Ischnura elegans, whereas androchromic females are recessive in Ischnura senegalensis. Here we report a case of interspecific hybridization between a gynochrome female of I. elegans and a male of I. senegalensis in the laboratory. We obtained 61 hybrid adult offspring, of which all 31 females were of gynochrome morph. DNA analyses of the hybrids confirmed that nuclear DNA sequences were derived from both parent species, whereas mitochondrial DNA sequences were maternally inherited. In the hybrids, the postocular spots of female heads, the shape of male appendages, and the color of female's cerci resembled those of I. elegans, whereas the size of abdominal blue spots was similar to that of I. senegalensis. The shape of prothorax and basal abdominal markings were intermediate in females. The larval developmental traits and the morphological changes in the final larval instar of the hybrids were similar to those of I. senegalensis. To our knowledge, this is the first report of hybrids between two damselfly species with different dominant‐recessive patterns of female color morphs.

Phylogeny of Macronematinae (Trichoptera: Hydropsychidae) based on molecular and morphological analyses

AbstractThe phylogenetic relationships among the members of Macronematinae (Trichoptera: Hydropsychidae) have long been debated and often revised. Our study based on a large subunit (28S) nuclear ribosomal DNA (D2 expansion fragment; 464 base pairs) of gene sequences found all genera to be monophyletic except for the genusPolymorphanisusWalker, which produced two clades in accordance with its species groups established using morphological characters. Phylogenetic reconstruction based on mitochondrial cytochrome oxidase I (658 base pairs) found the same monophyletic relationships, however, except for the generaPolymorphanisusandMacrostemumKolenati. The placement of the genusLeptonemaGuérin-Méneville as the basal clade in the subfamily was also ruled out, which is consistent with previous morphological findings. The monophyly of two previously recognised tribes (Macronematini and Polymorphanisini) was not supported. We also found that the generaCentromacronemaUlmer,BaliomorphaNeboiss, andMacronemaPictet are most closely related, having three head setal warts on the vertex of the head and unsegmented inferior appendages of male genitalia as their synapomorphies. Our study supported the hypothesis of close relationship of the generaAmphipsycheMcLachlan andProtomacronemaUlmer with the morphological synapomorphies: absence of a forewing discoidal cell and the presence of elevated head carinae on the larvae. Some notable evolutionary novelties in the structure of adults in various lineages of this subfamily include reduction of maxillae and labium, reduction and loss of a discoidal cell and evolution of ac-sccrossvein in each forewing, fusion of segments in each male inferior appendage, and evolution of dark colour patterns in the forewings. Also, the head setal warts, which are unique organs in Trichoptera, evolved towards a decreasing number in the subfamily. This first molecular phylogenetic study, covering most genera in Macronematinae and including an interpretation of 12 key evolutionary novelties, provides an important basis for resolving long-standing questions regarding phylogenetic relationships and classification of genera and species and helps lay a stronger foundation for inference of testable hypotheses about functional traits for species of this common and widespread subfamily.

A review of Dendrocephalus (Dendrocephalinus) (Crustacea: Anostraca) with the first records of male-male anostracan aggressive competition

We present a review of Dendrocephalus (Dendrocephalinus) with an updated diagnosis for the subgenus and a key to all known species. We provide new records of Dendrocephalus alachua, which was previously supposed to be extinct, and we describe a new species, Dendrocephalus proeliator sp. nov., which is separated from all other species based on the form of the male frontal appendage. Dendrocephalus proeliator sp. nov. appears to be morphologically intermediate between D. alachua and D. lithaca. In addition, we provide conservation assessments for all four species in the subgenus, according to IUCN Red List standards. We also report for two species the first known examples of direct male-male agonistic behaviour and competition for access to areas frequented by receptive females.

Female Wolf Spider, Schizocosa avida1, Vibration Receptor Responses to Male Courtship

Spiders are model systems for exploring proximate and ultimate questions about the evolution of vibratory signals. However, little is known of the sensory response characteristics of receivers, with one exception in Cupiennius spp. Spiders have vibration receptor organs called metatarsal organs on their legs. The organs contain aggregations of slit sensilla that respond to cuticular strain caused by substrate vibrations. Two neurons innervate each sensillum. How distal and proximal slit sensilla responded to male vibratory courtship components were shown. Neurons in isolated female legs were recorded extracellularly as a male courted nearby. Across 5 females, 9 slits were recorded in total. In one preparation, both distal and proximal slit responses to male vibratory courtship were captured. Altogether, distal slit responses captured primarily vibrations produced by male leg taps while proximal slit responses captured vibrations produced by male leg taps and drumming of pedipalps (male appendages used for sperm transfer). In addition, leg taps of variable amplitude elicited similar neural responses in all recorded slits. Compared to preparations in which female leg tarsi were immobilized, treatment preparations of movable female tarsi provided reliable neural responses to male courtship. Probing sensory processing features of females allows exploration into how male signals are encoded. Such work could inform how receiver sensory systems might influence the evolution of male signals.

Identification and analysis of odorant-binding protein genes from the wolf spider Pardosa pseudoannulata (Araneae: Lycosidae) based on its transcriptome

Pardosa pseudoannulata, one of the dominant predators in the paddy ecosystem, is a potential resource in the biological control of rice pests. Olfaction is crucial for insect behaviors such as feeding, mating and foraging, and odorant-binding proteins (OBPs) play a vital role in smell sensing, so we suppose that OBPs may be also important in spider behaviors. In this study, we identified two putative OBP genes and obtained their complete cDNA sequences. PpseOBP1 is 842 bp long and encodes a 188 amino acids protein. PpseOBP2 is 996 bp long and encodes a 213 amino acids protein. By our predictions and analysis, both of the PpseOBPs share some similar physicochemical and structural characteristics with insect classic OBP genes, but they do not strictly observe the cysteine pattern of OBPs in insects, in which there are three amino acid residues between the second and the third cysteine residues but 27 amino acid residues in P. pseudoannulata. Phylogenetic analysis showed that two PpseOBPs clustered together at the edge of the tree, indicating their distant relationship with insects. Regarding insects, PpseOBPs were most closely related to the Periplaneta americana OBP1. Both PpseOBPs are mainly expressed in appendages (pedipalps and legs). Expression of PpseOBP1 was significantly higher in female appendages than in male appendages, while PpseOBP2 showed an extreme expression level in male pedipalps. This is the first report of molecular characterization and expression patterns of OBPs in Lycosidae and will establish a foundation for better understanding of molecular mechanisms of P. pseudoannulata olfaction and facilitate its function in biological control.

Description of a new species of Nososticta Hagen (Odonata: Platycnemididae: Disparoneurinae) from Central Nicobar Islands, India.

A new damselfly species, Nososticta nancowra Rajeshkumar sp. nov. is described based on eight males and three females from Central Nicobar Islands, India. The diagnostic characteristics of the new species separating it from congeners are discussed. Morphological variations observed in the anal appendages of males and the pronotum of females are documented. A description of the new species using photographs and drawings is given. An identification key to species of the genus Nososticta occurring in Nicobar Islands is given for both sexes.

New species of the bristletail genus Allopsontus Silv. (Microcoryphia, Machilidae) from Southeastern Kazakhstan and Abkhazia

Two new bristletail species, Allopsontus (Anisoptinus) nigrus sp. n. from Southeastern Kazakhstan and A. (Kaplinilis) abkhazicus sp. n. from Abkhazia, are described. A. nigrus sp. n. differs from the only other known species of the subgenus Anisoptinus, A. borgustani Kaplin 2015, in a number of characters, namely, in more rounded paired ocelli, an extended apical segment of the labial palpus, a long articulated ovipositor, and in the absence of short appressed pigmented setulae on the maxillary and labial palpi in males. A. abkhazicus sp. n. can be easily distinguished from all the other known species of the subgenus Kaplinilis by the absence of short appressed pigmented setulae on the maxillary and labial palpi in the males, the absence of a projection with numerous spiniform setae on the male fore femur, and in shorter male genital appendages. The ocellus of the male and female is 2.8-2.9 times as wide as long; the length to width ratio of the eye is 0.9-1.0; the ovipositor consists of 60 divisions.

Redescription of Tripaphylus musteli (van Beneden, 1851) (Copepoda: Sphyriidae) and the relegation of Paeon Wilson, 1919 to synonymy with Tripaphylus Richiardi in Anonymous, 1878

Tripaphylus musteli (van Beneden, 1851) (Copepoda, Siphonostomatoida, Sphyriidae) is redescribed from an adult female collected from the branchial chamber of a starry smooth-hound, Mustelus asterias Cloquet (Carcharhiniformes, Triakidae), captured in the English Channel off Portland, UK. The new account of T. musteli is the first based on a complete adult female and highlighted the lack of a robust distinction separating Tripaphylus Richiardi, in Anonymous, 1878 and Paeon Wilson, 1919 prompting us to relegate Paeon to a junior subjective synonym of Tripaphylus. In the light of this synonymy the eight former species of Paeon are transferred to Tripaphylus as follows: T. ferox (Wilson, 1919) new combination, T. elongatus (Wilson, 1932) new combination, T. vassierei (Delamare Deboutteville & Nuñes-Ruivo, 1954) new combination, T. lobatus (Kirtisinghe, 1964) new combination, T. asymboli (Turner, Kyne & Bennett, 2003) new combination, T. versicolor (Wilson, 1919) new combination, T. australis (Kabata, 1993) new combination, and T. triakis (Castro Romero, 2001) new combination. Comparisons between terminology used in this report and that in the literature indicate that all transformed adult females of Tripaphylus probably possess a full complement of cephalic appendages and maxillipeds. All limbs, with the exception of the maxillae share a general morphological similarity to the corresponding appendages of conspecific males. The maxilla of the transformed adult female of Tripaphylus is a small digitiform protuberance associated with a swelling in some species.

Leptobasis linda sp. nov. from Ecuador (Odonata: Coenagrionidae).

Leptobasis linda is described from the Pacific lowlands of Ecuador. The coloration of mature individuals is superficially similar to the widespread L. vacillans, but structural and color characteristics differentiate L. linda from all congeneric species. The male caudal appendages and the female posterior margin of the prothorax are unique among Leptobasis.

Rediscovery and redescription of Coenagrion persicum (Lohmann 1993) with description of the female, and some notes on habitat selection (Odonata: Coenagrionidae).

Coenagrion persicum was described by Heinrich Lohmann in 1993 on the basis of a single male and two larvae captured in 1937 by E.W. Kaiser in Lorestãn Province (W-Iran). In June 2015 two of the authors (TS and DI) rediscovered individual-rich populations of this species in two Iranian provinces (Lorestãn and Esfahãn). We could confirm the structural differences of the male appendages between C. persicum and C. pulchellum based on a larger number of specimens than in the original description. The structural differences from C. pulchellum in females and their phenotypic variation pattern is described. Coenagrion persicum and C. pulchellum are also genetically distinct regarding two nDNA gene fragments: arginine methyltransferase (PRMT) and phosphoglucose isomerase (PGI). In contrast with C. pulchellum, C. persicum prefers small springs and running waters with rich herbal vegetation. Our faunistic data indicate that the species is present in the mountains between 1800 m and 2300 m a.s.l.. The species seems to be restricted to W-Iran, where it co-occurs with other rheophilic species.

Geometric morphometrics and molecular systematics of Xanthocnemis sobrina (McLachlan, 1873) (Odonata: Coenagrionidae) and comparison to its congeners.

The taxonomy of the damselfly genus Xanthocnemis is revised, with particular focus on populations inhabiting the North Island of New Zealand. Earlier studies revealed two species: X. sobrina, restricted to cool, shaded streams in kauri forests and other forested areas, and X. zealandica, a common species throughout New Zealand except the Chatham and subantarctic islands. A field study encompassing aquatic habitats throughout the whole North Island was carried out to establish the relationship between morphological variation (body size and various morphological traits over the entire body) observed by previous researchers with ecological conditions and/or geographical location. The main aim was to propose reliable diagnostic features that could be used in future studies. Morphological and molecular variation was assessed. Morphological examination included assigning landmarks for all body parts corresponding to the external morphological features that are usually used in Odonata taxonomy. Molecular analysis targeted fragments of the 28S and 16S rRNA genes. Congruence was sought between both types of data, statistical support for two morphological types previously described as different species and a maximum likelihood phylogenetic tree in conjunction with a pairwise genetic distance matrix constructed from the DNA sequences obtained from the sampled specimens. Geometric morphometrics revealed statistically significant differentiation between specimens identified as X. zealandica and X. sobrina for four traits: (1) dorsal view of the head for both sexes as well as male appendages from (2) dorsal, (3) ventral and (4) lateral views. Wings appeared different when analysed for males only. Molecular analysis, however, grouped all specimens into a single undifferentiated cluster with very low mean pairwise distance (<0.01) between them showing almost no variation at the molecular level among the sampled populations on the North Island. Therefore, an additional analysis of the mitochondrial cytochrome c-oxidase I gene was carried out comparing randomly selected North Island specimens to Xanthocnemis specimens targeted in other molecular studies (Nolan et al. 2007, Amaya-Perilla et al. 2014). The analysis of the COI gene confirmed that all North and South Island isolates of Xanthocnemis cluster together in a well-supported clade with pairwise identity >96% and ~93% pairwise identity with X. tuanuii sequences obtained from the Chatham Island specimens. A careful investigation of the thin plate spline deformations generated for the geometric morphometric landmarks showed that the significant variations in the appendages of the Xanthocnemis specimens appeared to be the result of size, rather than shape, differences. Therefore, X. sobrina is proposed as a synonym of X. zealandica. Recently Amaya-Perilla et al. (2014) synonymised X. sinclairi with X. zealandica and confirmed the status of the Chatham Island X. tuanuii as a distinct species. It is therefore proposed that the genus Xanthocnemis consists of two species only: zealandica occurring all over the North, South and Stewart Islands, and tuanuii, endemic to Chatham and Pitt islands. Considering several statistical tests involving body measurements and ecological variables recorded during the field study, as well as various discussion points from similar studies of other species of Odonata, two alternative hypotheses are proposed for future testing. The first hypothesis synonymises X. sobrina with X. zealandica and suggests a possible explanation for the evolution of the two morphological traits that have previously been considered diagnostic for these species. The second hypothesis suggests that as typical X. sobrina were not sampled during this study this could represent a species that is now extinct, unless future studies prove it otherwise.

Redescription of Two Krithid Species (Crustacea, Ostracoda) from the Sea of Japan, with a Comment on the Taxonomic Characters of Krithidae

Abstract. The marine podocopid ostracod Krithe antisawanensis is redescribed on the basis of its appendages and the carapaces of adult male and female specimens. The krithid species Parakrithella pseudadonta, which has only been previously described on the basis of the carapace morphology and male appendages excluding the copulatory organ, is redescribed here with its female appendages and male copulatory organ. Both krithid genera (Krithe and Parakrithella) have three articulated podomeres in the fifth and sixth limbs and an axe-head anterior vestibulum. Such characters are useful as diagnostic features for the family.

'A virtue beyond all medicine': The Hanged Man's Hand, Gallows Tradition and Healing in Eighteenth- and Nineteenth-century England.

From the eighteenth century through to the abolition of public executions in England in 1868, the touch of a freshly hanged man's hand was sought after to cure a variety of swellings, wens in particular. While the healing properties of corpse hands in general were acknowledged and experimented with in early modern medicine, the gallows cure achieved prominence during the second half of the eighteenth century. What was it about the hanged man's hand (and it always was a male appendage) that gave it such potency? While frequently denounced as a disgusting ‘superstition’ in the press, this popular medical practice was inadvertently legitimised and institutionalised by the authorities through changes in execution procedure.

Morphological variation in males of Dendrocephalus orientalis (Anostraca: Thamnocephalidae): Implications for species identification.

The genus Dendrocephalus Daday, 1908 consists of 17 described species. The morphology of the frontal appendage and the first pairs of thoracopods of males are the main characteristics used for species identification. The objectives of this study were to increase knowledge of the morphology of Dendrocephalus orientalis and to propose a new identification key for the species occurring in Brazil. Specimens were collected in temporary ponds within the conservation unit Monument Natural Grota do Angico (MNGA), which straddles the borders of the municipalities of Poço Redondo and Canindé de São Francisco, State of Sergipe, Brazil. During the study period, 560 male individuals of D. orientalis were sampled. The specimens showed variations in the number of spines on the eyes and the proximal surface of the male frontal appendage. On the eye, the number of spines ranged from 0 to 2; morphotypes with 1 and 2 spines predominated in the samples, with 50.5% and 46.2%, respectively. The number of spines on each eye was the same within each individual. On the proximal surface of the males frontal appendage the following numbers of spines on each arm were recorded: 1 and 1 (57.86%); 2 and 1 (30.89%); 2 and 2 (9.46%); 1 and 0 (1.25%); 3 and 2 (0.36%); 2 and 0 (0.18%). Based on these variations, we conclude that the number of spines on the eyes and the frontal appendage arms cannot be used as a diagnostic character for species identification. On the other hand, the first pairs of thoracopods and sub-branches 1V and 2A of the frontal appendage should be included in keys, since these structures did not show significant intraspecific morphological variation.

Visual Perception in the Brain of a Jumping Spider

Jumping spiders (Salticidae) are renowned for a behavioral repertoire that can seem more vertebrate, or even mammalian, than spider-like in character. This is made possible by a unique visual system that supports their stalking hunting style and elaborate mating rituals in which the bizarrely marked and colored appendages of males highlight their song-and-dance displays. Salticids perform these tasks with information from four pairs of functionally specialized eyes, providing a near 360° field of view and forward-looking spatial resolution surpassing that of all insects and even some mammals, processed by a brain roughly the size of a poppy seed. Salticid behavior, evolution, and ecology are well documented, but attempts to study the neurophysiological basis of their behavior had been thwarted by the pressurized nature of their internal body fluids, making typical physiological techniques infeasible and restricting all previous neural work in salticids to a few recordings from the eyes. We report the first survey of neurophysiological recordings from the brain of a jumping spider, Phidippus audax (Salticidae). The data include single-unit recordings in response to artificial and naturalistic visual stimuli. The salticid visual system is unique in that high-acuity and motion vision are processed by different pairs of eyes. We found nonlinear interactions between the principal and secondary eyes, which can be inferred from the emergence of spatiotemporal receptive fields. Ecologically relevant images, including prey-like objects such as flies, elicited bursts of excitation from single units.